Dromedary (Camelus dromedarius) and Bactrian camel (Camelus bactrianus) crossbreeding husbandry practices in Turkey and Kazakhstan: An in-depth ...

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Dromedary (Camelus dromedarius) and Bactrian camel (Camelus bactrianus) crossbreeding husbandry practices in Turkey and Kazakhstan: An in-depth ...
Dioli Pastoralism: Research, Policy and Practice         (2020) 10:6
https://doi.org/10.1186/s13570-020-0159-3
                                                                                                             Pastoralism: Research, Policy
                                                                                                                             and Practice

 REVIEW                                                                                                                                        Open Access

Dromedary (Camelus dromedarius) and
Bactrian camel (Camelus bactrianus)
crossbreeding husbandry practices in
Turkey and Kazakhstan: An in-depth review
Maurizio Dioli

  Abstract
  Hybridization between dromedaries and Bactrian camels has been carried out since ancient times in various areas
  of the Middle East and Central Asia. The primary force behind such practice was to obtain a better pack camel for
  trade caravans and military interventions. Today, camel hybridization is carried out systematically in only two areas:
  Turkey and Kazakhstan. In Turkey, camel cross-breeding is mainly done to obtain large animals for the annual camel
  wrestling competitions, while in Kazakhstan, cross-breeding is done to obtain camels that are more productive and
  better able to resist the harsh climate of Central Asia.
  Keywords: Arabian camel, One-humped camel, Dromedary, Camelus dromedarius, Camel, Two-humped camel, Bactrian
  camel, Camelus bactrianus, Hybridization

Introduction                                                                         indicated by archaeological remains (Mukhareva 2007;
The dromedary, also known as one-humped camel or                                     Potts 2005; Peters and Driesch 1997, Pigière and Henrotay
Arabian camel (Camelus dromedarius), and the Bactrian                                2012) and images of Bactrian camels on common objects
camel or two-humped camel or simply camel (Camelus                                   such as Roman coins (Fig. 2). The dromedary and Bactrian
bactrianus) are two distinctive and recognizable livestock                           camel, although taxonomically distinct species, are capable
species commonly found in Africa, the Middle East, and                               of interbreeding with each other. This produces a fertile
Asia. The two species occupy disparate geographic areas,                             F1 (first filial generation) hybrid that, thanks to the hybrid
with the dromedary found in the northern part of Africa,                             vigour or heterosis arising from the interaction between
the Middle East, part of Asia, and the Indian subcontinent,                          parental genomes, has a better growth rate and producti-
while the Bactrian camel is found only in inner, central                             vity than that of their pure-bred parents (Dzhumagulov
and east Asia: China, Mongolia, Kazakhstan, Kirghizstan,                             1980; Baimukanov 2002). More importantly, F1 hybrids are
Turkmenistan, and Afghanistan, northern areas of Iran,                               capable of withstanding colder, wetter climates and rough
India, Pakistan and up to eastern Turkey (Fig. 1). Present-                          terrains with greater ease. These peculiarities were powerful
day Bactrian camel distribution is much smaller than the                             incentives toward the development by various ethnic groups
area occupied by the dromedary, which is essentially lim-                            of structured cross-breeding husbandry practices between
ited to areas with a mean annual temperature not much                                dromedary and Bactrian camels. It is postulated that
over 21 °C (Mason 1979). This was not always the case.                               dromedary-Bactrian camel cross-breeding was initiated
since historically, certainly from first millennia BCE to the                        around the second century BCE, or even much earlier,
XVIII century CE, the distribution of the Bactrian camel                             particularly in areas where the distribution of the two species
was much wider, extending to include Russia, much of the                             overlapped or wherever female dromedaries were common
Middle East, Turkey, the Balkans, and Eastern Europe, as                             and Bactrian camels could be easily obtained: in various areas
                                                                                     of the Middle East, Turkey, from Iran, Iraq to southern
                                                                                     Arabia, and from Turkmenistan to northern Afghanistan
Correspondence: camel4ever@fastemail.us
Khartoum, Sudan
                                                                                     (Bulliet 1975, 2009; Potts 2004, 2005; Uerpmannh 1999).

                                      © The Author(s). 2020 Open Access This article is distributed under the terms of the Creative Commons Attribution 4.0
                                      International License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and
                                      reproduction in any medium, provided you give appropriate credit to the original author(s) and the source, provide a link to
                                      the Creative Commons license, and indicate if changes were made.
Dromedary (Camelus dromedarius) and Bactrian camel (Camelus bactrianus) crossbreeding husbandry practices in Turkey and Kazakhstan: An in-depth ...
Dioli Pastoralism: Research, Policy and Practice   (2020) 10:6                                                            Page 2 of 20

 Fig. 1 Map of approximate distribution and overlapping of Camelus dromedarius and Camelus bactrianus. www.maxpixel.net

Historical dromedary-Bactrian camel cross-                              documented repeatedly (Bulliet 2009; Tapper 1985,
breeding husbandry practices                                            2011) and is probably the reason why skeletal remains
The main driver of dromedary and Bactrian cross-                        of hybrid camels have been found in various archaeo-
breeding practices was the need to obtain strong pack                   logical sites where trade centres and military garrisons
camels capable of withstanding rough terrains and the                   were located. Such sites can be found all over Turkey
colder, wet climates of geographical areas with large trade             and northern, eastern, and central Europe (Pigière and
volumes and subjected to military conquests and cam-                    Henrotay 2012; Vukovic and Bogdanovic 2013; Çakırlar
paigns. These geographical areas included modern day                    and Berthon 2014; De Grossi Mazzorin 2006, 2010;
area of Turkey, Balkans, and northern and eastern Europe.               Bartosiewicz 1996; Bartosiewicz and Dirjec 2001; Galik
The exceptional loading capacity of hybrid camels is                    et al. 2015). The Ottoman army used hybrid camels ex-
                                                                        tensively because of their exceptional loading capacity:
                                                                        400–500 kg and ability to withstand cold and hilly ter-
                                                                        rains (Leese 1927; Burckhardt 1831; van Lennep 1870;
                                                                        Russell and Russell 1794). During the 1529 siege of
                                                                        Vienna by the Ottoman army, thousands of camels
                                                                        were used and scores ended up as spoils of war after
                                                                        the Ottoman defeat (Schimmer 1847). Such camels
                                                                        were probably the base for the image of the sixteenth-
                                                                        century engraving by Niklas St n of a “Turk with
                                                                        camel and a dromedary” depicting a Bactrian camel
                                                                        that, by showing two very small humps close to each
                                                                        other, must be described as a hybrid between a drom-
                                                                        edary and a Bactrian (Fig. 3). The description of an
                                                                        atrophic hump, anterior to the main hump, in adults
                                                                        and foetuses of the dromedaries bred at San Rossore,
                                                                        Italy, between the seventeenth and twentieth centuries
                                                                        (Lombardini 1879), is a further proof that camel
                                                                        hybridization was rather common in Europe (Dioli
                                                                        2014, 2018). Camel cross-breeding was carried out by
                                                                        mating the male Bactrian camel with the female drom-
                                                                        edary. Such an approach is confirmed by archaeological
 Fig. 2 Roman coin from the Roman province of Arabia, Bostra Syria
                                                                        findings that have identified the dromedary as the ma-
 Decapolis AD 98–117. www.catawiki.com
                                                                        ternal species used in the cross-breeding (Uerpmannh
Dromedary (Camelus dromedarius) and Bactrian camel (Camelus bactrianus) crossbreeding husbandry practices in Turkey and Kazakhstan: An in-depth ...
Dioli Pastoralism: Research, Policy and Practice   (2020) 10:6                                                                   Page 3 of 20

 Fig. 3 Engraving of a “Turk with camel and a dromedary” from Niklas Stӧn dated 1530 showing the single indented hump typical of dromedary/
 Bactrian hybrids (courtesy Herzogliches Museum Landenmuseum-Gotha, Germany)

1999). This cross-breeding practice was the most com-                   relatively common (Adamova and Rogers 2004) (Fig. 4).
mon method reported by several authors (Leonard                         In all historical illustrations of “camel wrestling”, the
1894; Marsh 1856; Plassio 1912; Steel 1890) and men-                    animals represented are not pure-bred dromedaries or
tioned by various regional travellers (Burckhardt 1831;                 Bactrian camels but hybrid F1. While the presence of a
Olearius 1669). A visual confirmation that this was in-                 single hump excludes them as pure-bred Bactrian
deed the cross-breeding practice used is even visible in                camels, their hump is too large and elongated to be one
an image of the famous sixteenth-century illuminated                    of the pure-bred dromedaries. In addition, the abun-
manuscript from Iran, Haft Awrang, in which an illus-                   dant hairs all along the front side of the neck, the back
tration shows a male Bactrian camel together with a fe-                 and top of the head, and the forearms rule out such an-
male dromedary and between them their hybrid calf                       imals as dromedaries since these characteristics are ab-
(Simpson 1997: 92). The first-generation hybrid F1                      sent in pure-bred dromedaries while constantly present
camel’s loading capacity and ability to work in hilly ter-              in Bactrian camels. Wrestling camels in paintings are
rains and cold environments was common knowledge.                       shown foaming at the mouth and without the “dulla” or
It was mentioned in the report documenting the                          soft palate, an organ present in the male dromedary but
nineteenth-century importation of dromedaries into the                  absent in the Bactrian camel and hybrids.
USA (United States Government Report 1857). In that
report, Lieutenant Porter, who was responsible for the                  Modern dromedary-Bactrian camel hybrid
expedition, even suggested the importation of pure                      production
male Bactrian camels that could be bred with female                     Currently, camel hybridization is done in eastern
dromedaries to produce F1 hybrids (United States Gov-                   Turkey (Yilmaz and Ertugrul 2014) and more exten-
ernment Report 1857: 123). Further cross-breeding of                    sively and in more structured ways in the former
F1 hybrid camels among themselves was rarely carried                    Soviet Republics of Kazakhstan and neighbouring
out since the resulting progeny was smaller, weaker,                    Turkmenistan and Uzbekistan (Faye and Konuspayeva
and “extremely vicious” or unable to cope with inclem-                  2012; Baimukanov et al. 2019; Dzhumagulov 1980;
ent weather (Burckhardt 1831; Fraser 1825; van Lennep                   Ruchkina and Vakhitova 2008). In Turkey, the main pur-
1870; Leonard 1894; Marsh 1856; Stewart 1881; Steel                     pose of hybrid production is to obtain large camels for the
1890). Another historical use of male Bactrian and fe-                  annual “wrestling” events organized in the south-western
male dromedary F1 hybrids was in the so-called camel                    part of the country during winter (Çakırlar and Berthon
wrestling events. Multiple mediaeval miniature paint-                   2014; Yilmaz 2017; Yilmaz et al. 2015a, b, Yilmaz and
ings of Islamic art historians depicting camel wrestling                Ertugrul 2014) (Fig. 5). In Kazakhstan and neighbouring
from various geographic areas such as Afghanistan,                      countries, the focus of hybridization is to obtain camels
western India, and Iran suggest that such events were                   better at milk, wool, and meat production while at the
Dromedary (Camelus dromedarius) and Bactrian camel (Camelus bactrianus) crossbreeding husbandry practices in Turkey and Kazakhstan: An in-depth ...
Dioli Pastoralism: Research, Policy and Practice   (2020) 10:6                                                                       Page 4 of 20

 Fig. 4 Illustration from Abd al Samad: Two Fighting Camels, ca.1590. Wikimedia
 Commons. https://commons.wikimedia.org/wiki/File:Abd_al_Samad._Two_Fighting_Camels_ca._1590._Private_Collection.jpg

same time more resilient to the harsh climate of Central                  currently carried out in Turkey (Yilmaz and Ertuğrul
Asia (Imamura et al. 2017).                                               2015; Yilmaz et al. 2015a, b). The Turkish hybrid pro-
                                                                          duction method is the basic “two-breed” cross consisting
Hybrid production methods from Turkey                                     of mating a male Bactrian camel with a female dromed-
In Turkey, hybrid production methods aim at producing                     ary (Fig. 7). In Turkish, the common name of such F1
a first-generation hybrid F1, occasionally a second gener-                hybrid is tülü, more specifically besrek for male and
ation F2 (second filial generation) and more rarely a                     maya for female (Yilmaz et al. 2015a, b). These hy-
third-generation hybrid F3 (third filial generation). Fig-                brids are very similar in appearance to the dromed-
ure 6 is illustrates the various combinations of                          ary, although they possess a larger and more
dromedary-Bactrian camel cross-breeding practices                         elongated hump which starts from just behind the

 Fig. 5 Two male F1 hybrid camels wrestling at the 2018 Camel Wrestling Festival in Izmir, Turkey. Note the profuse foam at their mouth and the
 attendants ready to separate the camels once a victor has emerged. This will prevent injuries
Dromedary (Camelus dromedarius) and Bactrian camel (Camelus bactrianus) crossbreeding husbandry practices in Turkey and Kazakhstan: An in-depth ...
Dioli Pastoralism: Research, Policy and Practice   (2020) 10:6        Page 5 of 20

 Fig. 6 Dromedary-Bactrian camel cross-breeding practices in Turkey
Dromedary (Camelus dromedarius) and Bactrian camel (Camelus bactrianus) crossbreeding husbandry practices in Turkey and Kazakhstan: An in-depth ...
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 Fig. 7 A Bactrian camel male mating with a female dromedary. Note the position of the male front legs comfortably straddling the female and
 so conferring stability to the male position

shoulder and extends well over the lumbar vertebrae.                     crosses between F1 are not done since the resulting
The peculiar shape is named “almond hump” or                             progeny, named kukirdi, have substandard body de-
badem horguc in Turkish (Fig. 8). In addition to be-                     velopment (Dzhumagulov 1980) and reputedly poor
ing unusually elongate, the hump of an F1 hybrid in                      behavioural qualities to such an extent that these has
the anterior area has a small indentation or notch a                     been described as “a menace to the mother and the
few centimetres deep. This is clearly visible in well-                   herd” (Çakırlar and Berthon 2014). However, further
nourished hybrid camels with short hair or detectable                    hybridization is occasionally done using the “back-
by manual palpation in the case of hybrid camels                         cross” method in which the males and females of the
with a long coat. F1 hybrids possess also abundant                       first cross-breeding are mated to dromedary or
hair all along the front side of the neck, the top and                   Bactrian camels obtaining second-generation hybrids
back of the head, and the forearms. Further F2                           (F2). They are named tavsi, when the female F1

 Fig. 8 A Turkish F1 hybrid tülü showing the typically elongated hump named “almond hump” in Turkish
Dromedary (Camelus dromedarius) and Bactrian camel (Camelus bactrianus) crossbreeding husbandry practices in Turkey and Kazakhstan: An in-depth ...
Dioli Pastoralism: Research, Policy and Practice   (2020) 10:6                                                            Page 7 of 20

maya is bred with a male Bactrian camel, and teke,                       genetic influence, they are adapted to colder and hilly
when a male F1 tülü is bred with a female dromed-                        areas of Turkey such the Taurus mountains.
ary. They are not used in wrestling, but as working
animals, tavsi for colder areas and teke for the hotter                  Hybrid production methods from Kazakhstan
areas of southern Turkey. F1 maya may also be bred                       Kazakhstan, being the largest country in former Soviet
to a male dromedary producing an F2 hybrid camel                         Central Asia with a population of both dromedaries and
named yegen; however, this is only done occasionally                     Bactrian camels, has the most complex camel hybridization
since his performance qualities are not appreciated                      practices with well-established cross-breeding strategies de-
(Yilmaz et al. 2015a, b). Further cross-breeding is                      veloped by Russian and Kazakh scientists (Dzhumagu-
done to produce F3 animals by back-crossing a fe-                        lov 1980; Baimukanov 2002; Ruchkina and Vakhitova
male teke with a Bactrian camel; such hybrids are                        2008). There are many regional linguistic differences
called kertelez and because of the higher Bactrian                       and dialects; therefore, the terminology identifying

 Fig. 9 Basic cross-breeding methods adopted in Russia, Kazakhstan, and neighbouring countries
Dromedary (Camelus dromedarius) and Bactrian camel (Camelus bactrianus) crossbreeding husbandry practices in Turkey and Kazakhstan: An in-depth ...
Dioli Pastoralism: Research, Policy and Practice   (2020) 10:6                                                                    Page 8 of 20

hybrids is complex (Bulliet 1975; Sala and Kartaeva                      do not differ much from each other. Both have a single
2017; Tapper 1985) and prone to confusion and                            very large hump extending from the shoulder to the lum-
inconsistencies on the correct terms for specific hy-                    bar area with a distinct notch, a few centimetres deep, on
brids (Imamura et al. 2016). This article follows the                    the front of the hump. These hybrids have also more hair
latest hybrid etymology adopted by Baimukanov                            than the pure-bred dromedary and in the same body areas
et al. (2019). Two main cross-breeding methods are                       of Bactrian camels: neck, head, hump, and forearms. As in
adopted, both initially based on a two-breed cross                       the case of horse cross-breeds, the phenotype of the
followed by various stages of back-crossing and are                      mother is slightly more obvious (Lin et al. 2016) and so
illustrated in Fig. 9. The so-called “Turkmen Hybridization              iner-maya hybrids tend to look more like a dromedary
Method” in which the first generation (F1) of camel                      while nar-maya inherit more Bactrian camel characteris-
hybrids is produced crossing a male Bactrian camel                       tics, particularly neck and forearm hair (Ruchkina and
with female dromedary results in a hybrid camel                          Vakhitova 2008).
named iner-maya for female and iner for male. The                           Male and female F1 hybrids are not bred among them-
“Kazakh Hybridization Method” in which a male                            selves since with such breeding the advantages of heterosis,
dromedary is crossed with a female Bactrian camel                        hybrid vigour, disappear. Furthermore, their progeny, jarbal
results in a F1 hybrid female camel named nar-maya                       or jarbay or dzharbai, have poor body development with
and a male named nar (Baimukanov et al. 2018,                            deformed chest and joints and an unsound temperament
2019) (Fig. 10). This method is possible because both                    (Baimukanov 2002; Dzhumagulov 1980; Ruchkina and
the Kazakh breed of dromedary and the Turkmen                            Vakhitova 2008). All further crossings of F1 hybrids are
dromedary or arvana (Fig. 11), has a large and long-                     therefore based on a back-crossing system: female F1 are
bodied conformation that allows the male to over-                        mated with one of their original species, while male F1 are
come the obstacle of the female Bactrian camel rear                      usually castrated (Ruchkina and Vakhitova 2008). The
hump when mating. Both dromedary and Bactrian                            back-crossing of nar-maya (F1) females with male Bactrian
camel mate in a crouched position; however, when a                       camels produces a group of F2 hybrids named kospak,
female Bactrian camel is mated with a male dromed-                       whose females will subsequently be mated with Turkmen
ary, her rear hump forces a short-bodied mating male                     male dromedaries to produce a group of F3 hybrid camels
dromedary to assume a very erect position which is                       named kez-nar. Iner-maya (F1) females back-crossing with
unstable. In such circumstances, the male often slides                   male Turkmen dromedaries produce a group of F2 hybrids
down sideways away from the female and is unable to                      named kurt whose females will later on be mated with male
complete the mating. F1 hybrids, iner-maya and nar-                      Kazakh Bactrian camels to produce a group of F3 camel
maya, or bir-tygan in Kyzylorda region of Kazakhstan,                    hybrids named kurt-nar (Figs. 12 and 13).

 Fig. 10 A dromedary male mating with a Bactrian camel female. The rear hump of the female and the short-bodied conformation of the male is
 forcing the male to assume a very erect mating position. In such conditions, the mating dromedary front legs are unable to reach the ground
 and the male loses balance and slide down on the female side and is unable to complete the mating
Dromedary (Camelus dromedarius) and Bactrian camel (Camelus bactrianus) crossbreeding husbandry practices in Turkey and Kazakhstan: An in-depth ...
Dioli Pastoralism: Research, Policy and Practice     (2020) 10:6                                                               Page 9 of 20

 Fig. 11 An adult dromedary male stud of the Arvana breed. This dromedary breed is long-bodied and large with a big elongated hump and
 dense brown coat. The breed is tolerant to cold wet conditions and has good milk production of 10–15 l daily. Man height, 1.85 m

  The “Turkmen Hybridization Method” and the                           Modern variations of the Turkmen Hybridization
“Kazakh Hybridization Method” have been known at                       Method
least for several decades (Dzhumagulov 1980); how-                     Figure 14 summarizes the various modifications done by
ever, renewed interest in Kazakhstan toward achieving                  Kazakh scientists to the Turkmen Hybridization
a more profitable utilization of the dromedaries and                   Method. As in the original hybridization scheme, the
Bactrian camels resulted in the development of more                    iner-maya, obtained by crossing a female dromedary
complex hybridization schemes.                                         with a male Bactrian camel, is back-crossed with a male

 Fig. 12 Hybrid camels: a kospak 3, b, c kez-nar, and d kurt
Dromedary (Camelus dromedarius) and Bactrian camel (Camelus bactrianus) crossbreeding husbandry practices in Turkey and Kazakhstan: An in-depth ...
Dioli Pastoralism: Research, Policy and Practice   (2020) 10:6                                                             Page 10 of 20

 Fig. 13 Hybrid camels: a iner-maya, b nar-maya, c kospak 1, and d kospak 2

dromedary of Turkmen breed to give birth to an F2                         2 produce an F4 hybrid named kurt-nar 2
hybrid named kurt. After this, different cross-breeding                   (Kazakhstan Patent Database 2009a).
strategies are implemented. The first F2, kurt or kurt 1,                   A three-breed cross can be carried out crossing an
is again back-crossed with a Turkmen dromedary to                         F4 kurt-nar 2 hybrid with a Kazakh dromedary pro-
produce an F3 hybrid named kurt 2, or sapaldryks, and                     ducing a new type of F5 hybrid called baytur
this then back-crossed again with a Turkmen dromedary                     (Kazakhstan Patent Database 2010c). A three-breed
to produce an F4 hybrid (fourth filial generation) named                  cross is also done crossing a female F2 kurt 1 with
kurt 3. This hybrid finally is back-crossed yet again to                  the Kazakh dromedary male to obtain a new type of
produce an F5 hybrid (fifth filial generation) named kurt                 F3 hybrid named bai-nar (Kazakhstan Patent Data-
4. This last hybrid, with 96.8% Turkmen dromedary                         base 2005). Bai-nar may occasionally be crossed with
genes and only 3.12% Bactrian camel genes, is then de-                    the new Kazakh dromedary or a pure-bred Bactrian
veloped further by breeding among itself and creating a                   camel, generating hybrids that are reported to reach
new breed, the so-called Kazakh dromedary (Baimuka-                       maturity at an earlier age. One can also cross an F3
nov 2019; Kazakhstan Patent Database 2003). Phenotyp-                     kurt 2 again with a Kazakh dromedary male to ob-
ically, this new breed is indistinguishable from a                        tain a new type of F4 hybrid named bekdas. Finally,
dromedary since the three successive back-crossings                       bekdas can be back-crossed with a Kazakh dromed-
with the Turkmen dromedary have stabilized the shape                      ary producing a new type of F5 hybrid named
of their single hump and removed the presence of the                      bekdas-nar (Kazakhstan Patent Database 2010d).
typical Bactrian body hair on the neck, forearms, and                       Within the Turkmen Hybridization Method, further
head (Fig. 15). However, the milk of Kazakh dromedary                     utilization of a “three-breed cross-breeding” using the new
has a higher fat content percentage than the milk of the                  breed of Kazakh dromedary has led to the creation of new
Turkmen dromedary with an average fat content of 4.5%                     hybrids. Figure 16 illustrates the various steps. It begins by
(Baimukanov et al. 2019) compared to the dromedary mean                   crossing a male of the new Kazakh dromedary with a fe-
fat content % of 3.52 ± 0.74% (Zhao et al. 2015). F3 kurt 2               male F1 iner-maya to produce a new F2 hybrid named bay-
can also be bred among themselves producing an F4                         shin. This will then be back-crossed with a pure male
commercial hybrid simply known as “hybrid”. How-                          Bactrian camel generating a new F3 hybrid named bay-
ever, female kurt hybrids are often back-crossed with                     kazhy-nar. Females of this new hybrid will be further back-
pure Bactrian camels. In such cases, F2 kurt 1 give                       crossed with a male Turkmen dromedary producing a new
birth to an F3 hybrid named kurt-nar 1 while F3 kurt                      F4 hybrid named ardas which will be back-crossed again
Dioli Pastoralism: Research, Policy and Practice     (2020) 10:6                                                                       Page 11 of 20

 Fig. 14 Modern variations of the Turkmen Hybridization Method leading to the production of new hybrids: bai-nar; bekdas and bekdas-nar; baytur;
 kurt-nar; kurt 1, 2, and 3; and “Kazakh dromedary”
Dioli Pastoralism: Research, Policy and Practice       (2020) 10:6                                                                         Page 12 of 20

 Fig. 15 Typical phenotypic characters of Bactrian camel: (a) two humps with their bases separated by 30–40 cm or more of flat area; (b) abundant
 forearms hair; (c) abundant hair localized on the front margins of the neck, from the throat area to the neck base; and (d) abundant hair on the top of
 the head between the ears. These areas of hair growth are absent in pure dromedaries while these are always present in varying degree in hybrids

with a Kazakh dromedary to produce a new F5 hybrid                             genetic make-up of 12.5% and a dromedary heritage of
named sannak (Baimukanov et al. 2016, 2017b, 2019).                            87.5% made up by the Turkmen dromedary breed
                                                                               (37.5%) and the new Kazakh dromedary breed (50%)
Modern variations of the Kazakh Hybridization                                  (Kazakhstan Patent Database 2009b). F2 arada-kurt may
Method                                                                         also be used to develop a line of hybrids named Dostik
The Kazakh Hybridization Method has been further                               by back-crossing repeatedly with dromedaries of the
modified by Kazakh scientists. Two main breeding plans                         Turkmen breed. The mating generates an F3 hybrid
have been developed, one involving crossing the nar-                           named Dostik 2 and subsequently an F4 hybrid Dostik 3.
maya with the newly developed breed of Kazakh drom-                            These latest hybrids are bred among each other and gen-
edary and another one using the Turkmen breed of                               erate an F5 hybrid simply named type Dostik 3 (Baimu-
dromedary. Figure 17 schematically shows the first ap-                         kanov et al. 2013; Kazakhstan Patent Database 2014).
proach. It is a three-way cross using the newly developed                        The second variation of the Kazakh Hybridization
breed Kazakh dromedary. The F1 nar-maya is back-                               Method is shown in Fig. 18 (Baimanukov 2019, Kazakh
crossed with a male of Kazakh dromedary to produce a                           Patent Database 2008a, 2009c, 2010a, b). With this
new F2 hybrid named aydaramir-arada. This hybrid will                          breeding method, the F1 nar-maya is back-crossed to a
be then back-crossed to a Bactrian camel to produce a                          male Bactrian camel to produce an F2 hybrid named
new F3 hybrid named aydaramir-nar that will be again                           kospak 1 or bal-kospak. This hybrid is then further back-
back-crossed to a dromedary of the Turkmen breed to                            crossed with a Bactrian male to obtain a new F3 hybrid
produce a new F4 hybrid named aydaramir-kurt. This                             named kopsak 2 or myrza kospak, and again to obtain a
latest hybrid will finally be back-crossed to the new                          new F4 hybrid named kospak 3 or nar-kospak
breed of Kazakh dromedary obtaining a new F5 hybrid                            (Kazakhstan Patent Database 2008b). All the hybrid kos-
named aydaramir (Baimukanov et al. 2019). The second                           paks can be back-crossed with Turkmen dromedaries to
hybridization strategy consists of crossing the F1 nar-                        produce various types of hybrids named kez-nars. F2
maya with a male dromedary of the Turkmen breed to                             kospak 1 produces F3 kez-nar 1, F3 kospak 2 produces
obtain an F2 hybrid arada-kurt (also named Dostik 1).                          an F4 kez-nar 2, and F4 kospak 3 produces an F5 kez-
This hybrid will be crossed with the new breed of Kaz-                         nar 3 (Kazakhstan Patent Database 2008a).
akh dromedary obtaining a new F3 hybrid named                                    An important addition to this cross-breeding scheme
arada-nar. Such hybrids are further developed and                              consists of introducing a “three-breed cross-breeding”
allowed to breed “in itself” ultimately producing an F4                        strategy crossing the F2 hybrid kospak with the new breed
hybrid that is considered a completely new camel breed                         of Kazakh dromedary and by doing so generating a com-
named arada. The new breed has a Bactrian camel                                pletely new F3 hybrid named baidara (Kazakhstan Patent
Dioli Pastoralism: Research, Policy and Practice   (2020) 10:6                                                                  Page 13 of 20

 Fig. 16 Turkmen Hybridization Method utilization of a “three-breed cross-breeding” using the new breed of Kazakh dromedary to produce new
 hybrids: bayshin, baykazhy, ardas, and sannak
Dioli Pastoralism: Research, Policy and Practice       (2020) 10:6                                                                           Page 14 of 20

 Fig. 17 Modern variations of the Kazakh Hybridization Method leading to the production of new hybrids: dromedary type Dostik 3, arada, and aydaramir
Dioli Pastoralism: Research, Policy and Practice   (2020) 10:6                                                         Page 15 of 20

 Fig. 18 Kazakh Hybridization Method variations leading to the production of new hybrids—bereket-nar and kospak 1, 2, and 3—and
 utilization of a “three-breed cross-breeding” using the new breed of Kazakh dromedary to produce the new hybrids: baidara
 and baidasbeck
Dioli Pastoralism: Research, Policy and Practice   (2020) 10:6                                                                     Page 16 of 20

Database 2009c). Baidara may occasionally be crossed                      and aidaramyr have a higher live weight than the
with the new Kazakh dromedary or a pure-bred Bactrian                     dromedary and the Bactrian camel. The work of
camel as this seems to increase early reproductive matur-                 Russian and Kazakh researchers (Baimukanov et al.
ity. The Kazakh dromedary is also utilized by crossing it                 2016, 2017a, b, c, 2018, 2019; Dzhumagulov 1980) has
with a female F5 kez-nar 3 to produce a new F6 hybrid                     established the following general rules concerning
(sixth filial generation) named baidasbeck (Kazakhstan Pa-                camel hybridization practices:
tent Database 2010b). Recently, a variation of this scheme
has been added by utilizing the larger breed of Bactrian                      Breeding hybrid males is not recommended (except
camel: Kalmyk breed (Fig. 19). When crossing a female F3                        F5 kurt 4, F4 Dostik 3, F3 arada-nar).
kospak with a male Bactrian camel of Kalmyk breed, a                          F1 hybrids are 5–12% larger in body size than each
valuable new F3 hybrid is obtained named bereket-kospak,                       of their own pure-bred parents, the dromedary and
which produces a new F4 hybrid named bereket-nar when                          Bactrian camel, and with better working ability and
crossed with a male Turkmen dromedary (Kazakhstan Pa-                          higher endurance. F1 hybrids obtained with the
tent Database 2010a).                                                          Kazakh Hybridization Methods are larger than F1
                                                                               obtained with the Turkmen Hybridization Method.
                                                                               The positive effects of heterosis are greater from the
Overview of main established hybrid advantages                                 age of 1 year onward.
The main objective of dromedary and Bactrian camel                            All hybrids have greater productive longevity than
cross-breeding practices developed by Kazakh and Rus-                          dromedary and Bactrian camels with F1 nar-maya
sian scientists has been to develop a camel breed with                         and F3 kez-nar capable of 7.3 and 6.8 lactations in
higher milk production capabilities than the Bactrian                          their lifetime respectively.
camel as well as the ability to survive the harsh climate                     When F1 hybrids breed among themselves, the
of Central Asia essentially in a free-range situation. The                     positive effects of heterosis are completely absent
climatic conditions of such regions are characterized by                       and the resulting F2 hybrid, named jarbay, is an
hot summers and snowbound winters with severe sub-                             unfit animal.
zero temperatures. Numerous hybrids with such qual-                           Hybrid calves have a higher birth weight
ities have been developed. Of particular interest are                          (F1 mean birth weight, 45.4 ± 0.842.1 kg) than
bainar, baykazhy, aydaramir-arada, aydaramir-nar,                              the calves of dromedaries (mean birth weight,
baidasbeck, and baidar hybrids which have a higher                             34.5 ± 0.09 kg; Nagy and Juhász 2019) and
milk production than the Turkmen dromedary. Mean-                              of Bactrian camels (mean birth weight,
while the F2 hybrids: bayshin, and aidaramyr-nar, all                          34.55 ± 7.17 kg; Zhao et al. 2000). Their growth
F4 hybrids (excluding kurt 3) and F5 hybrids sannak,                           rate is also higher.

 Fig. 19 Bactrian camel stud of the Kalmyk breed. This Bactrian camel breed is massive, capable to reach a live body weight of more than 1200
 kg. The breed can produce 6–9 kg of fine wool and has a daily milk production of 2.5–3.5 l. Man height, 1.85 m
Dioli Pastoralism: Research, Policy and Practice    (2020) 10:6                                                                      Page 17 of 20

   Higher percentages of dromedary genes in a                             Conclusions and recommendations
      hybrid increases overall milk production due to:                     Camel hybridization may play an important role in enhan-
      longer and more abundant lactation; improves                         cing camel productivity in all areas of Central Asia that
      udder size and conformation including better quarters                presently keep only Bactrian camels because of the inability
      symmetry and development; increased teats spacing,                   of dromedaries to survive in the Central Asian climate. In-
      size, and downward direction. However, this also                     deed, trends toward increasing camel milk production util-
      reduces fat percentage in milk, reduces wool                         izing hybrids are already detectable in areas of Kazakhstan
      production and early maturity as well as adaptability                (Imamura et al. 2017). In many areas of Kazakhstan, drom-
      to colder climates.                                                  edaries, Bactrian camels, and their hybrids are free-ranging
     Higher percentage of Bactrian camel genes in a                       and naturally intermingling (Fig. 20). In such conditions,
      hybrid increases wool production, live weight, and                   unsupervised mating may occur and go unrecorded. produ-
      fat percentage in milk but decreases lactation length                cing animals with an unknown dromedary/Bactrian camel
      and absolute milk yield. The length of pregnancy                     genetic make-up. All such hybrids, while morphologically
      also increases.                                                      identical to pure-bred dromedaries, still retain Bactrian
     All hybrids have a gestation length shorter than the                 camel genes that may influence a hybrid’s productivity. The
      one of the Bactrian camel (mean 442.5 ± 5.1 days) but                identification of hybrid camels based on phenotypic charac-
      longer than that of the Turkmen dromedary (mean                      ters is unreliable, particularly for later stage hybrids such as
      425 ± 3.9) and Kazakh dromedary (mean 417.2 ± 3.1                    F4-F6 (Table 1 and Fig. 15). Rigorous pedigree recording,
      days). Dromedary genes shorten the duration of                       combined with permanent camel identification, are there-
      pregnancy. F4 hybrids have a gestation length closest                fore better tools to clearly identify individual hybrids and
      to the Turkmen and Kazakh dromedary.                                 the exact genetic contributions of dromedary and Bactrian
     All hybrids have a lower wool production than that                   camel to their genetic make-up. The increasing availability
      of Bactrian camels                                                   of advanced techniques for genetic and parentage testing
     All hybrids have a lower fat percentage in milk than                 will allow a better understanding of the genetics of drom-
      that of Bactrian camels (mean 5.34 ± 0.60) but                       edary, Bactrian camel, and their hybrids, possibly permit-
      higher than that of pure-bred dromedaries (mean                      ting the detection of valuable genetic traits related to
      3.52 ± 0.74) (Zhao et al. 2015)                                      increased productivity or adaptability (Burger et al. 2019).

 Fig. 20 a, b Hybrid camels of unknown generation possibly F4-F5. c Hybrid camels: left to right, bereket-kospak and iner-may. d Hybrid camels:
 left to right, kez-nar, kospak, and kez-nar 3
Dioli Pastoralism: Research, Policy and Practice          (2020) 10:6                                                                         Page 18 of 20

Table 1 List of dromedary-Bactrian camel hybrids mentioning their phenotypic Bactrian characters
Hybrids                                                                 Bactrian camel Camelus bactrianus common phenotypic traits present in hybrids
                                                                        heritage %
Hybrid name             F   Parents                                                    Hump area      Areas with Bactrian hair
                            ♀                  ♂                                                         Forearms           Neck           Top head
Nar-maya                F1 Bactrian            Dromedary                50               2               In all camel hybrids, the typical Bactrian camel
                                                                                                         hair grown in these three body areas is always
Iner-maya               F1 Dromed.             Bactrian                 50                               present.
Kospak 1                F2 Nar-maya            Bactrian                 75               6               However, hair presence and extent are noticeable
                                                                                                         only in fall, winter, and spring seasons. In the
Aydaramir-arada         F2 Nar-maya            New Kazakh dromed. 25+                    1               summer/hot season, hair may be completely
Kurt 1                  F2 Iner-maya           Dromedary                25               3               shed rendering the aspect of these areas as in
                                                                                                         the dromedary
Bayshin                 F2 Iner-maya           New Kazakh dromed. 25+                    1
Arada-kurt (Dostik 1)   F2 Nar-maya            Dromedary                25
Aydaramir-nar           F3 Aydaramir-arada Bactrian                     62.5+            2
Kospak 2                F3 Kospak 1            Bactrian                 87.5             6
Bereket-kospak          F3 Kospak 1            Kalmyk Bactrian          87.5
Kurt 2                  F3 Kurt 1              Dromedary                12.5             2
Bai-nar                 F3 Kurt 1              New Kazakh dromed 12.5+
Kurt-nar 1              F3 Kurt 1              Bactrian                 62.5
Kez-nar 1               F3 Kospak 1            Dromedary                37.5             4
Baidara                 F3 Kospak 1            New Kazakh dromed. 37.5+                  1?
Baykazhy-nar            F3 Bayshin             Bactrian                 62.5             1
Arada-nar               F3 Arada-kurt          New Kazakh dromed. 12.5+
Dostik 2 type           F3 Arada-kurt          Dromedary                12.5
Dostik 3 type           F4 Dostik 2 type       Dromedary                6.25
Kospak 3                F4 Kospak 2            Dromedary                93.75            7
Aydaramir-kurt          F4 Aydaramir-nar       Dromedary                31.25+           ?
Kurt-nar 2              F4 Kurt 2              Bactrian                 56.25            2
“Hybrid F4”             F4 Kurt 2              Kurt 2                   12.5             1
Kurt 3                  F4 Kurt 2              Dromedary                6.25
Bekdas                  F4 Kurt 2              New Kazakh dromed. 6.25+
Kez-nar 2               F4 Kospak 2            Dromedary                43.75            4
Bereket-nar             F4 Bereket-kospak      Dromedary                43.75            3
Arada                   F4 Arada-nar           Arada-nar                12.5             1
Ardas                   F4 Baykazhy-nar        Dromedary                31.25
Baytur                  F5 Kurt-nar 2          New Kazakh dromed. 28.12+                 1
Aydaramir               F5 Aydaramir-kurt      New Kazakh dromed. 15.62+
Bekdas-nar              F5 Bekdas              New Kazakh dromed. 6.25+
New Kazakh dromed. F5 Kurt 4                   Kurt 4                   3.12
Kez-nar 3               F5 Kospak 3            Dromedary                46.87            3
Sannak                  F5 Ardas               New Kazakh dromed. 15.62+                 1
Baidasbeck              F6 Kez-nar 3           New Kazakh dromed. 23.43+
Explanatory notes: Hump area—8, Bactrian humps; 7, two well-developed humps closer than in the Bactrian but separate at their base; 6, two well-developed
humps with their bases touching each other; 5, one large hump with two symmetric small humps at the apex; 4, one large hump with two asymmetric small
humps at the apex, the rear hump bigger than the front one; 3, one large hump with two visible mounds at the apex; 2, one large hump with a clear depression
several centimetres deep on the front top margin; 1, dromedary hump. Bactrian heritage: +, to indicate that New Kazakh Dromedary has still 3.12% of Bactrian
camel genes
Dioli Pastoralism: Research, Policy and Practice             (2020) 10:6                                                                                   Page 19 of 20

Abbreviation                                                                          Bartosiewicz, L. 1996. Camels in antiquity: The Hungarian connection. Antiquity
F1: First filial generation                                                                70: 447–453.
                                                                                      Bartosiewicz, L., and J. Dirjec. 2001. Camels in antiquity: Roman period finds from
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My personal thanks to His Highness Sheikh Mohammed bin Hamad bin                      Bulliet, R.W. 1975. The camel and the wheel. Cambridge, Mass: Harvard University Press.
Mohammed Al Sharqi, the Crown Prince of Fujairah, for generously                      Bulliet, R.W. 2009. Of Turks and camels. In Cotton, climate, and camels in early
promoting and supporting this camel research. Thanks to Mr. Mohamed Bin                    Islamic Iran: A moment in world history, 96–126. Columbia University Press.
Ati, Mr. Adam, and Dr. Logman Abdalla Ahmed for their invaluable help.                Burckhardt, J.L. 1831. Notes on the Bedouins and Wahabys: Collected during his travels
Sincere thanks also to Dr. Barry Greville-Eyres for editing the manuscript’s               in the east / by late John Lewis Burckhardt. Vol. II. London: Colburn & Bentley.
English.                                                                              Burger, P.A., E. Ciani, and B. Faye. 2019. Old World camels in a modern world – a
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Author’s contributions                                                                     Genetic. https://doi.org/10.1111/age.12858.
The manuscript conceptualization, data collection, analysis, and writing were         Çakırlar, C., and R. Berthon. 2014. Caravans, camel wrestling and cowrie shells:
done by the author. The author read and approved the final manuscript.                     Towards a social zooarchaeology of camel hybridization in Anatolia and
                                                                                           adjacent regions. Anthropozoologica 49 (2): 237–252.
Funding                                                                               De Grossi Mazzorin, J. 2006. Cammelli nell’antichita: le presenze in Italia. In Studi
Not applicable.                                                                            di Archeozoologia in onore di Alfredo Riedel, ed. B. Sala and U. Tecchiati, 231–
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Availability of data and materials                                                    De Grossi Mazzorin, J. 2010. VIE DEGLI ANIMALI, VIE DEGLI UOMINI. Transumanza e
All data generated or analysed during this study are included in this published            altri spostamenti di animali nellʼEuropa tardoantica e medievale, 91–106. Italy:
article.                                                                                   Edipuglia s.r.l.
                                                                                      Dioli, M. 2014. The missing hump: A clarification on the reported two-humped
Ethics approval and consent to participate                                                 stage in the embryonic development of the dromedary (Camelus
Not applicable.                                                                            dromedarius). Journal of Camel Practice and Research 21 (2): 121–125.
                                                                                      Dioli, M. 2018. Camels in Europe: A brief history of camel hybrids and camel
Consent for publication                                                                    breeding in Europe over the last 1000 years. In Second Selçuk-Ephesus
Open access                                                                                Symposium on culture of camel-dealing and camel wrestling, 18-20 January
                                                                                           2018, Selçuk, Izmir, ed. A. Koç and Ö.Ü.H. Erdogan, vol. 2, 8–11. Izmir.
Competing interests                                                                   Dzhumagulov, I. K. 1980. Interspecific hybridization of camels (Characterization of
The author declares that he has no competing interests.                                    biological and productive qualities of camels, methods of breeding hybrids)
                                                                                           [Межвидовая гибридизация верблюдов. (Характеристика биологических
Received: 12 November 2019 Accepted: 4 January 2020                                        и продуктивных качеств верблюдов, методы разведения гибридов)],
                                                                                           Unpublished PhD dissertation, K. A. Timiryazev Moscow Agricultural
                                                                                           Academy, Moscow. [in Russian].
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